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ALEXANDRE F. BANNIKOV
(Borisyak Paleontological Institute of the Russian Academy of Sciences, Moscow)
A NEW PERCOID FISH PERCIFORMES
RELATED TO HENDRIXELLA
FROM THE EOCENE OF BOLCA, ITALY
ABSTRACT
e new genus and species of perciform fi sh Robertannia sorbiniorum gen. et sp. nov. is described from the Eocene locality
Pesciara of Bolca in northern Italy; it is shown to be a percoid related to the genus Hendrixella, also from the Pesciara of
Bolca. e new family Robertanniidae is established to accommodate Robertannia and Hendrixella. e sister-group rela-
tionship of this family among the Percoidei is unknown.
Key words: Perciformes, Percoidei, new taxa, Eocene, northern Italy, Bolca locality.
RIASSUNTO
Viene descritto Robertannia sorbiniorum gen. et sp. nov., un nuovo genere e specie di perciforme proveniente dalla località
eocenica della Pesciara di Bolca, Italia settentrionale; il percoideo presenta analogie con il genere Hendrixella, anch’esso pro-
veniente dalla Pesciara di Bolca. Viene istituita la nuova famiglia Robertanniidae per accogliere Robertannia and Hendrixella.
È sconosciuto il “sister-group” di questa famiglia tra i Percoidei.
Parole chiave: Perciformes, Percoidei, nuovo taxa, Eocene, Italia settentrionale, Bolca.
INTRODUCTION
e world-famous Eocene locality Monte Bolca (Italy) is exceptionally rich in marine
fi shes, which usually are very well preserved. e materials from Monte Bolca are widely
distributed among the world museums of natural history. e recently renewed emphasis on
excavations at the classic Pesciara site of the Monte Bolca locality, conducted by the Museo
Civico di Storia Naturale di Verona, has already brought forth many scientifi cally exciting new
materials of fossil coral-reef and lagoonal associated fi shes whose descriptions are beginning
to be published. Among these is the skeleton of a percoid fi sh found in 2009. e imprint is
the complete skeleton of a fi sh preserved in part and counterpart. is fi sh strongly resembles
the incertae sedis percoid Hendrixella grandei recently described from the Monte Bolca locality
(Bannikov and Carnevale, 2009). e new genus and species Robertannia sorbiniorum gen. et
. 8
sp. nov. is described below as a representative, along with Hendrixella Bannikov et Carnevale,
2009, of a new family of percoid fi shes, the Robertanniidae.
METHODS
Some details of the specimen examined were best seen when the specimen was moistened
with alcohol. e specimen was prepared by needle.
Interneural and interhaemal spaces are numbered based on the vertebra whose neural or
haemal spine forms the anterior border of the space, with the fi rst space being between the
fi rst and second neural or haemal spines (following Baldwin and Johnson, 1993; Bannikov
and Tyler, 1995; Tyler and Bannikov, 1997; etc.).
Abbreviations are as follows: Institutional: MCSNV – Museo Civico di Storia Naturale
di Verona; Anatomical: H – hypural; Ph – parhypural; PU – preural vertebra; SL – standard
length; U – ural vertebra.
SYSTEMATIC DESCRIPTION
Order Perciformes
Suborder Percoidei
Family Robertanniidae fam. nov.
D. Percoids with elongate body, moderate to rather large jaw teeth, lower jaw
articulation under orbit, preopercle not serrated, 6 or 7 branchiostegal rays, 10+14-16=24-26
vertebrae, most abdominal vertebrae bearing parapophyses, short vertebral spines, short ribs,
two series of intermuscular bones, unfused hypurals, 3 epurals, 2 supraneurals, two dorsal
fi ns with one or two rayless pterygiophores in-between, 7-9 slender and fl exible spines (one
supernumerary) in fi rst dorsal fi n, a spine and 8 soft rays in second dorsal fi n, anal fi n with
2 spines and 8 soft rays, anal fi n opposed to and nearly symmetrical with second dorsal fi n,
relatively narrow pectoral fi ns, pelvic fi ns inserted posterior to pectorals, relatively small and
forked caudal fi n with 17 principal rays, scales moderately large and weakly ctenoid.
T G. Robertannia gen. nov.
C. Type genus and Hendrixella Bannikov et Carnevale, 2009.
Genus Robertannia gen. nov.
D. Head moderate, ca. 31% SL; teeth relatively strong; 6 branchiostegal rays;
10+16=26 vertebrae; neural spines of abdominal vertebrae expanded; 7 spines in fi rst dorsal
fi n; two rayless pterygiophores between dorsal fi ns; base length of fi rst dorsal fi n equals to
distance between dorsal fi ns; pelvic fi ns inserted well posterior to pectorals.
T S. Robertannia sorbiniorum gen. et sp. nov., by monotypy and designation
herein.
() HENDRIXELLA , 9
E. e genus is named in honour of two stellar staff members of the Museo
Civico di Storia Naturale di Verona, Drs. Roberto Zorzin (curator) and Anna Vaccari (collec-
tion manager), in recognition of their continuing help in facilitating my research on Monte
Bolca fi shes; gender feminine.
Robertannia sorbiniorum sp. nov.
Figs. 1-3
D. at of the genus, of which it is presently the only known representative.
E. e patronym is in honour of the three fi ne paleoichthyologists of the Sor-
bini family in Verona and Pisa, my good friends and colleagues in the study of Italian fossil
fi shes: Prof. Lorenzo Sorbini and Drs. Margherita and Chiara Sorbini.
H. MCSNV IG VR 71980/71981, part and counterpart, complete skeleton, 47
mm SL; uppermost Lower or lowermost Middle Eocene (Medizza, 1975; Papazzoni and Tre-
visani, 2006), zone Discoaster sublodoensis; Monte Bolca locality, Pesciara cave site.
R S. None.
D. e body is elongate, with a moderately long caudal peduncle. e caudal
peduncle depth is 0.52 of the body depth. e head is moderately large, its length (tip of
snout to anterior edge of upper part of pectoral girdle) about 1.4 times greater than the body
depth. e head length is contained 3.24 times in SL. e maximum body depth is con-
tained 4.48 times in SL. e body seems to be somewhat compressed laterally.
- Head. e head depth is less than its length. e orbit is relatively small and placed
approximately in the middle of the head length. e mouth is terminal, with the lower jaw
articulation situated under the orbit. e neurocranium is relatively low and devoid of a
projecting supraoccipital crest. e frontals are relatively long and overhang the lateral eth-
moids. e mesethmoid is a well-ossifi ed block of bone. A narrow basisphenoid is present, as
evidenced by the holotypic counterpart MCSNV IG VR 71981. e infraorbital bones are
preserved indistinguishably. e upper jaw bones are greatly damaged. e premaxilla has a
thin ascending process separated from the articular process. e presence of a postmaxillary
process is unknown. e few preserved upper jaw teeth are relatively strong and conical; there
are faint indications on the presence of smaller teeth as well. e maxilla is narrow at its ar-
ticular head; the bone is expanded distally. e supramaxilla appears to be absent. e lower
jaw is relatively poorly preserved, with the limits of its bones scarcely distinguishable. e
dentary occupies most of the lower jaw; it seems to be moderately shallow near its symphysis.
e dentary is incised posteriorly for its articulation to the angulo-articular. ere are indica-
tions of a lateral line sensory canal in the lower portion of the dentary. e lower jaw teeth
are very poorly preserved, except for a single relatively strong tooth posteriorly on the dentary
of MCSNV IG VR 71981. Although the hyomandibula is not completely preserved, it is
evident that its shaft is inclined anteroventrally, and its head is relatively broad. e pterygoid
bones are too poorly preserved to be described. e quadrate is relatively small. Most of the
opercular bones are severely damaged. e preopercle seems to be rather wide and moderately
curved, with an apparently even posterior margin. No spines are evident on the posterior part
. 10
of the opercle; the bone is thickened along its anterior border. e ceratohyal has a distinct
slender anteroventral process to support the lower hypohyal. ere seems to be six sabre-like
branchiostegal rays. e urohyal is very low in its anterior region. Neither branchial bones
nor pharyngeal dentition are evident.
- Axial skeleton. ere are 26 vertebrae, ten abdominal and sixteen caudal, including the
urostyle. e vertebral column is almost straight. e vertebral centra are robust and slightly
elongate anteroposteriorly; most of them bear a longitudinal ridge on the lateral surface.
e fi rst vertebra is fore-shortened, whereas the centrum of the second vertebra is one of
the longest. e length of the caudal portion of the vertebral column is 1.45 times greater
than the length of the abdominal portion of the vertebral column. e vertebral spines are
slender and slightly curved. e neural spines of the abdominal vertebrae are expanded and
tapered to a point distally; those of the third and subsequent vertebrae are strongly inclined
posteriorly. e neural spines of the caudal vertebrae are slender and arise from the posterior
half of the centrum obliquely posterodorsally. e haemal spines are slender; those of the
anterior caudal vertebrae arise from the anterior half of the centrum. e haemal spines are
relatively strongly posteroventrally inclined. e haemal spine of the fi rst caudal vertebra has
a sigmoid-like curve. ere are indications that at least six posterior abdominal vertebrae bear
parapophyses. ere are eight pairs of pleural ribs, which are short, curved and slender; these
Fig. 1 – Robertannia sorbiniorum gen. et sp. nov., holotype: A – part, MCSNV IG VR 71980, B – counterpart,
MCSNV IG VR 71981; uppermost Lower or lowermost Middle Eocene, Monte Bolca locality in northern
Italy, Pesciara. Scale division: 1 mm.
() HENDRIXELLA , 11
Fig. 2 – Robertannia sorbiniorum gen. et sp. nov., reconstruction of the caudal skeleton based on the holotype
MCSNV IG VR 71980/71981. Scale bar = 2 mm.
. 12
are inclined posteriorly. e ribs of at least the anterior fi ve pairs bear slender intermuscular
bones. One more intermuscular bone is evident on MCSNV IG VR 71981, attached most
probably to the base of the neural spine of the fi rst vertebra. e latter intermuscular bone
seems to represent a diff erent series than those attached to the pleural ribs, as was shown for
Hendrixella grandei (Bannikov and Carnevale, 2009). In this taxon we regarded the intermus-
cular bones of the upper series as representing epineurals, whereas those of the lower series as
the homologs of the epicentrals of lower teleosts (Bannikov and Carnevale, 2009: 487). Our
interpretation contradicts to that of Patterson and Johnson (1995).
- Pectoral fi n and girdle. In MCSNV IG VR 71981 the posttemporal is relatively small
and forked. e elongate supracleithrum connects the posttemporal with the cleithrum and
obscures the fi rst vertebra laterally. e cleithrum is an elongate and relatively large bone situ-
ated below the fi rst and second vertebrae; the upper part of its length is curved forward. No
serrations are evident on the posterodorsal fl ange of the cleithrum. e coracoid is narrow
and wedge-shaped. e outlines of the scapula are unclear. e postcleithrum is a rib-like
bone extending obliquely from the posterodorsal fl ange of the cleithrum to the posterior por-
tion of the pelvic bone. e four pectoral radials are constricted in the middle and increase in
length below. e pectoral fi n is narrow; it seems to be relatively short. e total complement
of pectoral-fi n rays is unknown; about 10 rays are recognizable in MCSNV IG VR 71980.
e pectoral-fi n base is situated under the third to fourth vertebrae, approximately in the
middle of the distance between the vertebral column and the ventral profi le of the body.
- Pelvic fi n and girdle. e pelvic bones are only moderately large and wedge-like; each
bone is tightly attached medially to its opposite member. e anterior ends of the pelvic
bones seem to terminate under the pectoral radials and do not touch the cleithrum. It is not
known whether the position of the pelvic bones is natural or is the result of posterior displace-
ment post-mortem. e pelvic fi n is relatively short; it is inserted well behind the pectoral-fi n
base, under the end of the fi fth vertebra. ere are fi ve soft rays in each pelvic fi n, in addition
to the slender and apparently fl exible pelvic-fi n spine.
- Supraneurals and dorsal fi ns. ere are faint remains of two supraneurals (predorsal
bones); the fi rst supraneural precedes the neural spine of the second vertebra, and the second
lies in the second interneural space. ere are two dorsal fi ns divided by a distance which
exceeds the length of 4.5 opposite vertebrae. e fi rst dorsal fi n originates above the end of
Fig. 3 – Robertannia sorbiniorum gen. et sp. nov., reconstruction of the skeleton based on the holotype; scales
omitted.
() HENDRIXELLA , 13
the fi fth vertebra and terminates over the 10th vertebra. ere are seven exceptionally slender
and evidently fl exible dorsal-fi n spines in the fi rst dorsal fi n. e fi rst spine is supernumerary
on the fi rst dorsal-fi n pterygiophore, it is 1.25 times shorter than the second spine. e sec-
ond and third spines are approximately equal in length, whereas succeeding spines decrease
in length posteriorly in the series. e dorsal-fi n spines are close-set anteriorly, becoming
more widely spaced posteriorly. e fi rst pterygiophore of the dorsal fi n has anterior trian-
gular lamina and penetrates down into the second interneural space, contacting from above
the neural spine of the third vertebra. e fourth to sixth interneural spaces have the ventral
shafts of one pterygiophore present, whereas the third interneural space accommodates two
pterygiophores. e pterygiophores are relatively strongly inclined posteriorly; they bear lon-
gitudinal strengthening ridges in the proximal shafts. e medio-distal parts (concurrent to
the dorsal profi le of the body) of the pterygiophores become longer progressively from the
fi rst to the eighth. e seventh and eighth sequential pterygiophores are rayless; these are
situated between the dorsal fi ns.
e second dorsal fi n originates above the border between the fourth and fi fth caudal
vertebrae and terminates over the eighth caudal vertebra. ere is a weak spine and eight
soft rays in the second dorsal fi n. e fi rst soft ray is unbranched, whereas the other rays are
branched. e rays decrease in length posteriorly in the series. e length of the longest soft
rays equals the length of the longest spines of the fi rst dorsal fi n and exceeds the base length
of the second dorsal fi n. Few anterior pterygiophores of the second dorsal fi n penetrate down
into the interneural spaces, whereas the proximal shafts of the posterior ones do not reach the
tips of the neural spines. ere are a total of 10 second dorsal-fi n pterygiophores; therefore,
the last pterygiophore appears to be rayless. e medial pterygiophores seem to be fused with
the proximal ones.
- Anal fi n. e anal fi n originates below the fi fth caudal vertebra, being situated approxi-
mately under the second dorsal fi n. e anal fi n is only poorly preserved, with three ante-
rior elements being missing, except for their bases. It seems likely that there are two spines
and eight soft rays in the anal fi n. e fi rst anal-fi n spine is supernumerary. e length of
the longest anal-fi n rays is unknown. e anal-fi n pterygiophores are slender and relatively
strongly inclined posteriorly; these rapidly decrease in length posteriorly in the series. Most of
the anal-fi n pterygiophores are represented by imprints in the matrix rather than bony sub-
stance. e proximal ends of the anal-fi n pterygiophores do not reach the tips of the haemal
spines of the caudal vertebrae.
- Caudal fi n and skeleton. ere is a fracture through the posterior portion of the caudal
skeleton; therefore, the entire length of the hypurals is unknown. e terminal centrum is
composed of the fusion of PU1, U1 and U2 (Fig. 2). e hypurals, parhypural and haemal
spine of PU2 seem to be autogenous. e parhypural has a developed parhypurapophysis.
e neural and haemal spines of PU3 are longer than those of the preceding vertebra. It is un-
clear whether the haemal spine of PU3 is autogenous or fused with the centrum. e neural
spine of PU2 is a short crest. ere are three epurals; the fi rst of these is the longest, whereas
the others are progressively shorter. ere is at least one uroneural, which forms the stegural.
e caudal fi n is relatively small and forked. ere are 17 principal rays in the caudal fi n, 15
of which are branched. e number of procurrent rays is less certain; there are at least eight
rays both above and below.
- Squamation. in and moderately large scales cover the entire body and head. Most of
scales exhibit multiple (up to 10) delicate basal radiating grooves (radii); there are also circuli
on the scale surface and minute tubercles on the apical fi eld. Since the scales overlap each
. 14
other, their ctenii mostly are obscured, although delicate ctenii are visible in some places. e
anterior portion of the lateral line is observable as a slightly arched line above the tips of the
neural spines of the middle abdominal vertebrae.
- Measurements. Measurements as a percent of the 47 mm SL of the holotype are as fol-
lows:
Head length from tip of snout to posterior border of opercle: 31
Maximum body depth: 21
Depth of caudal peduncle: 11
Distance between tip of snout and fi rst dorsal fi n: 41.5
Distance between tip of snout and second dorsal fi n: 68
Distance between tip of snout and anal fi n: 70.5
Distance between pelvic fi n and anal fi n: 28
Length of base of fi rst dorsal fi n: 13.6
Length of base of second dorsal fi n: 9.5
Distance between dorsal fi ns: 13.6
Length of base of anal fi n: 9.4
Length of longest spine of fi rst dorsal fi n: 13.5
Length of longest soft ray of second dorsal fi n: 13.5
Length of longest ray of caudal fi n: 23
Length of longest spine of pelvic fi n: 9.8
Preorbital distance: 10.6
Horizontal diameter of orbit: 6.8
Length of lower jaw: ca. 18.7
R. Robertannia sorbiniorum gen. et sp. nov. defi nitely was a predatory fi sh. ere is
pray (partly articulated bones of the skeleton of a small unidentifi able fi sh) in the gut of the
holotype. e relatively strong dentition also indicates a predatory mode of life of the species
under consideration.
DISCUSSION
In many respect Robertannia sorbiniorum gen. et sp. nov. resembles the recently described
Hendrixella grandei from the same Monte Bolca locality (Bannikov and Carnevale, 2009).
ese two taxa have elongate bodies and share such characters as the preopercle with an even
posterior border, most of the abdominal vertebrae with parapophyses, short vertebral spines,
short ribs, two series of intermuscular bones, similar caudal skeletons, slender and fl exible
fi n spines, two widely divided dorsal fi ns, single supernumerary spine in the fi rst dorsal fi n,
a spine and 8 soft rays in the second dorsal fi n, 2 spines and 8 soft rays in the anal fi n, the
anal fi n opposed to and nearly symmetrical with the second dorsal fi n, the pelvic fi ns inserted
posterior to the narrow pectorals, the relatively small and forked caudal fi n with 17 principal
rays, and moderately large and weakly ctenoid scales. Such strong similarity justifi es the at-
tribution of Robertannia gen. nov. and Hendrixella to a single taxon of the familial level. Since
Hendrixella was regarded as a percoid genus incertae sedis (Bannikov and Carnevale, 2009),
the new percoid family Robertanniidae is established herein to accommodate Robertannia
gen. nov. and Hendrixella.
Robertannia sorbiniorum gen. et sp. nov. diff ers from Hendrixella grandei by having a larger
() HENDRIXELLA , 15
head and stronger dentition, apparently six branchiostegal rays (vs. seven in H. grandei), 16
caudal vertebrae (vs. 14 in H. grandei), expanded neural spines of the abdominal vertebrae
(vs. slender neural spines in H. grandei), seven spines in the fi rst dorsal fi n (vs. nine dorsal-fi n
spines in H. grandei), two rayless pterygiophores between the dorsal fi ns (vs. a single rayless
pterygiophore in H. grandei), stronger divided dorsal fi ns and more posterior insertion of the
pelvic fi ns in relation to the pectorals. ese diff erences justify the separate generic status of
Robertannia gen. nov. and Hendrixella.
e analysis of the distribution of selected derived features already performed for Hendrix-
ella (Bannikov and Carnevale, 2009) did not provide any convincing evidence that would
unite it to any previously known percoid family or incertae sedis genera. Similarly, the rela-
tionships of Robertanniidae fam. nov. within the Percoidei are rather diffi cult to defi ne.
e new discovery adds one more extinct family of percoid fi shes to the Eocene fi sh
fauna of Monte Bolca, in addition to the Ductoridae (see Blot, 1969), Exelliidae (see Blot,
1969; Bannikov and Tyler, 1994), Quasimullidae (see Bannikov, 1999), Carangodidae (see
Blot, 1969), Eocottidae (see Bannikov, 2004), and more than 10 incertae sedis fossil percoid
genera.
ACKNOWLEDGEMENTS
I am very grateful to Dr. Alessandra Aspes, former director of the Museo Civico di Storia
Naturale di Verona, for providing funding for my travel in 2010 to Verona to engage in this
and other studies. I also thank Dr. Roberto Zorzin and Dr. Anna Vaccari at the MCSNV for
their continuing help in facilitating my research on the fi shes of Monte Bolca, and Mrs. Bruna
Burato for her aid with the literature on Bolca fi shes. e author’s research was supported by
the Russian Foundation for Basic Research, grants nos. 08-05-00654 and 09-05-00170. Dr.
James C. Tyler of the Smithsonian Institution kindly revised the manuscript and improved
the English. Mr. Francesco Sorbini (Verona) kindly made the photograph for Figure 1.
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ALEXANDRE F. BANNIKOV
Borisyak Paleontological Institute of the Russian Academy of Sciences,
Profsoyuznaya 123, Moscow 117997, Russia
e-mail: aban@paleo.ru
